504:
484:
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452:, and high light-collecting ability, the ocelli are superbly adapted for measuring changes in the perceived brightness of the external world as an insect rolls or pitches around its body axis during flight. Locusts and dragonflies in tethered flight have been observed to try and "correct" their flight posture based on changes in light. Other theories of ocellar function have ranged from roles as light adaptors or global excitatory organs to polarization sensors and
259:
463:
fields of both the photoreceptors and the second-order neurons can be quite restricted. Further research has demonstrated these eyes not only resolve spatial details of the world, but also perceive motion. Second-order neurons in the dragonfly median ocellus respond more strongly to upwards-moving bars and gratings than to downwards-moving bars and gratings, but this effect is only present when
391:
the head, while a central (median) ocellus is directed forwards. In some terrestrial insects (e.g. some ants and cockroaches), the median ocellus is absent. The sideways-facing ocelli can be called "lateral ocelli", referring to their direction and position in the triplet, however this is not to be confused with the stemmata of some insect larvae, which are also known as lateral ocelli.
332:, respectively), most insects have dorsal ocelli while stemmata are found in the larvae of some insect orders. Despite the shared name, they are structurally and functionally very different. Simple eyes of other animals may also be referred to as ocelli, but again the structure and anatomy of these eyes is quite distinct from those of insect dorsal ocelli.
56:
567:, a family of sawflies) are only "simple" in that they represent immature or embryonic forms of the compound eyes of the adult. They can possess a considerable degree of acuity and sensitivity, and can detect polarized light. They may be optimized for light sensitivity, as opposed to detailed image formation. In the
271:
291:
a reflector at the back of the eyes. The light-sensitive part of the receptor cells is next to this, so they get direct and reflected light. In hunting or jumping spiders, for example, a forward-facing pair possesses the best resolution (and even some telescopic ability) to help spot prey from a distance.
390:
The number, forms, and functions of the dorsal ocelli vary markedly throughout insect orders. They tend to be larger and more strongly expressed in flying insects (particularly bees, wasps, dragonflies and locusts) where they are typically found as a triplet. Two ocelli are directed to either side of
290:
Spiders do not have compound eyes, but instead have several pairs of simple eyes with each pair adapted for a specific task or tasks. The principal and secondary eyes in spiders are arranged in four, or occasionally fewer, pairs. Only the principal eyes have moveable retinas. The secondary eyes have
462:
Recent studies have shown the ocelli of some insects (most notably the dragonfly, but also some wasps) are capable of "form vision" similar to camera eyes, as the ocellar lens forms an image within, or close to, the photoreceptor layer. In dragonflies it has been demonstrated that the receptive
432:
of the lens, as well as high convergence ratios and synaptic gains (amplification of photoreceptor signals), the ocelli are generally considered to be far more sensitive to light than the compound eyes. Additionally, given the relatively simple neural arrangement of the eye (small number of
406:. The number of photoreceptors also varies widely, but may number in the hundreds or thousands for well-developed ocelli. In bees, locusts, and dragonflies, the lens is strongly curved; while in cockroaches it is flat. Locusts possess vitreous humour while blowflies and dragonflies do not.
575:
role is their lateral position on the head; ocelli, that in other ways resemble stemmata, tend to be borne in sites median to the compound eyes, or nearly so. Among some researchers, this distinction has led to the use of the term "lateral ocelli" for stemmata.
467:
light is used in the stimulus; when ultraviolet light is absent, no directional response is observed. Dragonfly ocelli are especially highly developed and specialised visual organs, which may support the exceptional acrobatic abilities of these animals.
424:
These two factors have led to the conclusion that, with some exceptions in predatry insects, the ocelli are incapable of perceiving proper images and are thus solely suitable for light-metering functions. Given the large aperture and low
1366:
Müller, CHG; Rosenberg, J; Richter, S; Meyer-Rochow, VB (2003). "The compound eye of
Scutigera coleoptrata (Linnaeus, 1758) (Chilopoda; Notostigmophora): an ultrastructural re-investigation that adds support to the Mandibulata concept".
173:
have unambiguously compound eyes consisting of multiple lenses (up to tens of thousands), but achieve an effect similar to that of a camera eye, in that each ommatidium lens focuses light onto a number of neighbouring retinulae.
475:. Designers of these craft face many of the same challenges that insects face in maintaining stability in a three-dimensional world. Engineers are increasingly taking inspiration from insects to overcome these challenges.
513:
species shown here, is typical of the order
Lepidoptera. The head of the larva bears more than one pair of stemmata, all of which are set low down and are far more widely placed than the mouthparts.
1589:
Markus
Friedrich (2006). "Ancient mechanisms of visual sense organ development based on comparison of the gene networks controlling larval eye, ocellus, and compound eye specification in
1355:. 3rd US-European Competition and Workshop on Micro Air Vehicle Systems (MAV07) & European Micro Air Vehicle Conference and Flight Competition (EMAV2007). Vol. 1. pp. 17–21.
683:
1535:
1139:
448:
One common theory of ocellar function in flying insects holds that they are used to assist in maintaining flight stability. Given their underfocused nature, wide
545:, rarely have any eyes other than stemmata at any stage of their lives (exceptions include the large and well-developed compound eyes of the house centipedes,
1709:
266:'s main ocelli (center pair) are very acute. The outer pair are "secondary eyes" and other pairs of secondary eyes are on the sides and top of its head.
495:
An example of a sawfly larva. It has just a single pair of stemmata, and they are set higher on its head than the position of stemmata on the heads of
503:
387:(mantises). These ocelli coexist with compound eyes; thus, most insects possess two anatomically separate and functionally different visual pathways.
445:(often the largest diameter neurons in the animal's nervous system), the ocelli are typically considered to be "faster" than the compound eyes.
1094:
Eric J. Warrant, Almut Kelber, Rita Wallén & William T. Wcislo (December 2006). "Ocellar optics in nocturnal and diurnal bees and wasps".
691:
1893:
1650:
944:
889:
1784:
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are both expressed in the compound eye but not the simple eye, no reported 'developmental' genes are uniquely expressed in the simple eye.
104:
The structure of an animal's eye is determined by the environment in which it lives, and the behavioural tasks it must fulfill to survive.
1668:
1257:
987:
766:
Meyer-Rochow, V.B.; Reid, W.A. (1993). "Cephalic structures in the
Antarctic nemertine Parborlasia corrugatus - are they really eyes?".
2012:
1043:
1959:
824:
491:
1702:
1676:
1063:
335:
Dorsal ocelli are light-sensitive organs found on the dorsal surface or frontal surface of the head of many insects, including
483:
552:
Behind each lens of a typical functional stemma lies a single cluster of photoreceptor cells, termed a retinula. The lens is
1080:
1096:
402:). The ocellar lens may be strongly curved or flat. The photoreceptor layer may also be separated from the lens by a clear
1210:
817:
1255:
Richard Berry, Joshua van Kleef & Gert Stange (May 2007). "The mapping of visual space by dragonfly lateral ocelli".
1206:"A spatiotemporal white noise analysis of photoreceptor responses to UV and green light in the dragonfly median ocellus"
1551:
1306:
456:
420:
Dorsal ocelli ubiquitously have massive convergence ratios from first-order (photoreceptor) to second-order neurons.
417:
of the lens is not typically sufficient to form an image on the photoreceptor layer, essentially it is out of focus.
2017:
1695:
928:
1888:
1640:
579:
157:(a retina-like cluster of photoreceptor cells), including the ommatidia of most insects and the central eyes of
472:
137:). By other criteria, the presence of a complex retina distinguishes the vertebrate camera eye from the simple
35:
101:", which have multiple lenses. They are not necessarily simple in the sense of being uncomplicated or basic.
1155:"Form vision in the insect dorsal ocelli: an anatomical and optical analysis of the dragonfly median ocellus"
409:
Two somewhat unusual features of ocelli are particularly notable and generally common between insect orders.
845:
Zieger, V.; Meyer-Rochow, V.B. (2008). "Understanding the
Cephalic Eyes of Pulmonate Gastropods: A Review".
521:
larvae bear no other form of eyes until they enter their final stage of growth. Adults of several orders of
108:
differ widely in the habitats in which they live, as well as their visual requirements for finding food or
2078:
1498:
129:
and are sometimes considered "simple" because a single lens collects and focuses an entire image onto the
2022:
1981:
1976:
1954:
1529:
1133:
600:
A number of genetic pathways are responsible for the occurrence and positioning of the ocelli. The gene
295:
spiders' eyes are very sensitive in low light levels and are large to capture more light, equivalent to
123:
is flexible, and must be interpreted in proper context; for example, the eyes of most large animals are
2002:
1934:
1883:
1602:
1353:
Design concepts for a novel attitude sensor for Micro Air
Vehicles, based on dragonfly ocellar vision
1105:
644:
227:
also have ocelli, either at the tips or bases of their tentacles. Some other gastropods, such as the
199:
1024:"Contribution of compound eyes and ocelli to steering of locusts in flight: I. Behavioural analysis"
571:
stage, such stemmata develop into fully fledged compound eyes. One feature offering a clue to their
1402:
Meyer-Rochow, Victor Benno (1974). "Structure and function of the larval eye of the sawfly Perga".
147:
which make up compound eyes. Additionally, not all invertebrate ocelli and ommatidium have simple
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1490:"The orthodenticle gene encodes a novel homeo domain protein involved in the development of the
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1742:
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Adriana D. Briscoe & Lars
Chittka (2001). "The evolution of color vision in insects".
1159:
403:
961:
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1023:
204:– which have randomly distributed pigment, and which have no other structure (such as a
1939:
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324:(eye), and literally means "little eye". In insects, two distinct ocellus types exist:
263:
258:
209:
1439:"P. J. Gullan and P. S. Cranston: The insects: an outline of entomology (4th edition)"
2067:
1929:
1848:
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1801:
1762:
1724:
1415:
779:
639:
601:
553:
449:
438:
283:
253:
213:
212:). The apparent "eye color" in these animals is red or black. Certain groups such as
113:
1470:
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866:
30:"Ocellus" redirects here. For the light-sensitive structure in dinoflagellates, see
2073:
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Moth larva about to moult; the new stemmata are visible behind the old head capsule
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109:
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have more complex eyes, including some with a distinct retina, lens, and cornea.
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or an optical arrangement which has a single lens without the sort of elaborate
42:
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is allelic to ocelliless, a mutation that stops ocelli from being produced. In
2027:
1924:
1772:
1614:
1485:
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Although stemmata are simple eyes, some kinds (such as those of the larvae of
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372:
232:
228:
162:
143:
94:
55:
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also have stemmata, and never develop compound eyes at all. Examples include
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with seemingly simple eyes also emulate retinal vision in various ways. Many
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1204:
Joshua van Kleef, Andrew
Charles James & Gert Stange (October 2005).
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195:
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61:
31:
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1971:
1944:
1039:
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434:
360:
356:
328:(top-most) ocelli, and lateral ocelli (often referred to as ocelli and
985:
Martin Wilson (1978). "The functional organisation of locust ocelli".
626:) promotes the expression of orthodenticle and possibly eyes absent (
517:
Stemmata (singular stemma) are a class of simple eyes. Many kinds of
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352:
205:
170:
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134:
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90:
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1997:
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269:
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54:
556:, and the body of the stemma has a vitreous or crystalline core.
471:
Research on the ocelli is of high interest to designers of small
1300:
Joshua van Kleef, Richard Berry & Gert Stange (March 2008).
1153:
Richard P. Berry, Gert Stange & Eric J. Warrant (May 2007).
568:
526:
348:
275:
224:
1691:
1484:
R. Finkelstein, D. Smouse, T. M. Capaci, A. C. Spradling &
1793:
344:
340:
125:
86:
1351:
Gert Stange, R. Berry & J. van Kleef (September 2007).
715:"Temporal properties of the lens eyes of the box jellyfish
441:), as well as the extremely large diameter of some ocellar
97:. These eyes are called "simple" to distinguish them from "
591:) with stemmata incompletely aggregated into compound eyes
1302:"Directional selectivity in the simple eye of an insect"
114:
an enormous variety of eye types are found in arthropods
311:
The term "ocellus" (plural ocelli) is derived from the
48:"Ocelli" redirects here. For the eye-like marking, see
608:, the rhodopsin Rh2 is only expressed in simple eyes.
1675:. Ent 425 (General Entomology) course tutorial.
905:
630:) and as such is essential for simple eye formation.
41:"Ocellation" redirects here. Not to be confused with
929:"Evolution of eye structure and arthropod phylogeny"
803:
1990:
1902:
1874:
1792:
1723:
931:. In Stefan Koenemann & Ronald Jenner (eds.).
1059:"An ocellar dorsal light response in a dragonfly"
1584:
1582:
235:have ocelli that allow light to penetrate their
1703:
1494:nervous system and ocellar visual structures"
713:O'Connor M, Nilsson DE, Garm A (March 2010).
394:A dorsal ocellus consists of a lens element (
8:
1534:: CS1 maint: multiple names: authors list (
1138:: CS1 maint: multiple names: authors list (
116:to overcome visual problems or limitations.
316:
282:ocelli (centre), and the upper part of its
1710:
1696:
1688:
1057:Gert Stange & Jonathon Howard (1979).
1639:Warrant, Eric; Nilsson, Dan-Eric (2006).
1511:
1327:
1231:
1172:
742:
65:with two compound eyes and three ocelli (
922:
920:
112:, and avoiding predators. Consequently,
675:
1527:
1131:
1894:Evolution of color vision in primates
1595:Arthropod Structure & Development
1097:Arthropod Structure & Development
933:Crustacea and Arthropod Relationships
909:L.Koch: a fisheye lens in a spider".
884:. The Darwin Press, Inc. p. 25.
231:, have much more sophisticated eyes.
7:
1258:Journal of Comparative Physiology A
988:Journal of Comparative Physiology A
935:. Crustacean Issues. Vol. 16.
927:C. Bitsch & J. Bitsch (2005).
882:Coral Reefs: Cities under the seas
622:Epidermal growth factor receptor (
25:
805:. Vol. 6. 2007. p. 790.
398:) and a layer of photoreceptors (
1783:
1437:Wilson, Robert J. (2010-09-19).
1083:from the original on 2007-12-17.
1046:from the original on 2007-12-25.
962:"How many eyes does a bee have?"
911:Proceedings of the Royal Society
825:University of California, Irvine
615:at least) the genes eyeless and
1677:North Carolina State University
1667:Meyer, John R. (5 March 2006).
1064:Journal of Experimental Biology
1028:Journal of Experimental Biology
847:American Malacological Bulletin
1645:. Cambridge University Press.
1443:Journal of Insect Conservation
1320:10.1523/JNEUROSCI.5556-07.2008
818:"Photoreceptors of cnidarians"
1:
1565:10.1146/annurev.ento.46.1.471
1211:Journal of General Physiology
329:
151:. Many have various forms of
1671:. Department of Entomology.
1416:10.1016/0022-1910(74)90087-0
1404:Journal of Insect Physiology
1174:10.1016/j.visres.2007.01.019
960:Brindley, Hal (2023-03-07).
780:10.1016/0040-8166(93)90072-S
18:Simple eyes in invertebrates
1822:Simple eye in invertebrates
1552:Annual Review of Entomology
1307:The Journal of Neuroscience
880:Murphy, Richard C. (2002).
198:have the simplest "eyes" –
2095:
2018:Infrared sensing in snakes
1022:Charles P. Taylor (1981).
251:
47:
40:
29:
1889:Evolution of color vision
1781:
1615:10.1016/j.asd.2006.08.010
1455:10.1007/s10841-010-9351-x
1381:10.1007/s00435-003-0085-0
1271:10.1007/s00359-006-0204-8
1118:10.1016/j.asd.2006.08.012
816:Martin, Vicki J. (2002).
735:10.1007/s00359-010-0506-8
301:rufous net-casting spider
243:Simple eyes in arthropods
165:and some other predatory
27:Simple eye without retina
563:and especially those of
507:The larva of one of the
473:unmanned aerial vehicles
36:Ocellus (disambiguation)
1499:Genes & Development
801:"Eye (invertebrate)".
592:
514:
500:
488:
317:
287:
267:
70:
34:. For other uses, see
2023:Monocular deprivation
1982:Underwater camouflage
1977:Structural coloration
1955:Disruptive coloration
1224:10.1085/jgp.200509319
717:Tripedalia cystophora
582:
506:
494:
486:
437:between detector and
273:
261:
58:
2003:Blindness in animals
1935:Counter-illumination
1884:Evolution of the eye
1513:10.1101/gad.4.9.1516
1077:10.1242/jeb.83.1.351
939:. pp. 185–214.
859:10.4003/006.026.0206
684:"Catalog - Mendeley"
645:Evolution of the eye
93:that occurs in most
81:(sometimes called a
1642:Invertebrate Vision
1607:2006ArtSD..35..357F
1110:2006ArtSD..35..293W
723:J. Comp. Physiol. A
201:pigment spot ocelli
178:Ocelli or eye spots
1950:Deimatic behaviour
1040:10.1242/jeb.93.1.1
1001:10.1007/BF00661380
593:
515:
501:
489:
288:
268:
71:
2061:
2060:
2053:Visual perception
2048:Underwater vision
2013:Feature detection
2008:Eyespot apparatus
1967:Eyespot (mimicry)
1915:Animal coloration
1718:Vision in animals
1652:978-0-521-83088-1
1314:(11): 2845–2855.
1167:(10): 1394–1409.
946:978-0-8493-3498-6
891:978-0-87850-138-0
650:Eyespot apparatus
50:Eyespot (mimicry)
16:(Redirected from
2086:
2033:Palpebral (bone)
1866:Schizochroal eye
1787:
1712:
1705:
1698:
1689:
1684:
1679:. Archived from
1669:"Photoreceptors"
1656:
1627:
1626:
1586:
1577:
1576:
1546:
1540:
1539:
1533:
1525:
1515:
1506:(9): 1516–1527.
1481:
1475:
1474:
1434:
1428:
1427:
1410:(8): 1565–1591.
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907:Dinopis Subrufus
902:
896:
895:
877:
871:
870:
842:
836:
835:
833:
827:. Archived from
822:
813:
807:
806:
798:
792:
791:
763:
757:
756:
746:
710:
704:
703:
701:
699:
694:on 24 March 2012
690:. Archived from
688:www.mendeley.com
680:
596:Genetic controls
415:refractive power
322:
286:(left and right)
133:(analogous to a
119:Use of the term
21:
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1633:Further reading
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913:(196): 198–222.
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768:Tissue and Cell
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541:, such as some
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404:vitreous humour
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163:Jumping spiders
85:) is a form of
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1661:External links
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519:holometabolous
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450:fields of view
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264:jumping spider
252:Main article:
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1861:Parietal eye
1821:
1807:Compound eye
1681:the original
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692:the original
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660:Parietal eye
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110:conspecifics
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2038:Pseudopupil
1920:Aposematism
1839:Mollusc eye
1559:: 471–510.
1034:(1): 1–18.
655:Mollusc eye
585:Scolopendra
573:ontogenetic
561:Lepidoptera
531:springtails
465:ultraviolet
369:damselflies
365:dragonflies
337:Hymenoptera
278:with three
248:Spider eyes
233:Giant clams
196:ribbonworms
126:camera eyes
95:vertebrates
83:pigment pit
43:Oscillation
2068:Categories
2028:Ommatidium
1960:coincident
1925:Camouflage
1903:Coloration
1844:cephalopod
1738:Chameleons
1591:Drosophila
1492:Drosophila
1486:N Perrimon
971:2023-08-30
671:References
613:Drosophila
611:While (in
606:Drosophila
539:Arthropoda
510:Acherontia
457:entrainers
373:Orthoptera
274:Head of a
229:Strombidae
121:simple eye
106:Arthropods
75:simple eye
2043:Rhopalium
1876:Evolution
1849:gastropod
1817:Eye shine
1812:Eagle eye
1743:Dinosaurs
1463:1366-638X
937:CRC Press
617:dachshund
589:Chilopoda
587:species (
547:Scutigera
543:Myriapoda
535:Thysanura
454:circadian
400:rod cells
359:(flies),
293:Nocturnal
192:flatworms
188:sea stars
184:jellyfish
144:ommatidia
1910:Albinism
1623:18089081
1573:11112177
1488:(1990).
1471:28082252
1338:18337415
1287:25806901
1279:17273849
1242:16260838
1191:14590003
1183:17368709
1126:18089077
1081:Archived
1044:Archived
867:86083580
788:18621228
753:20131056
665:Ocelloid
634:See also
554:biconvex
523:hexapods
479:Stemmata
439:effector
435:synapses
385:Mantodea
353:sawflies
330:stemmata
154:retinula
62:Polistes
59:Head of
32:Ocelloid
1972:Mimicry
1945:Crypsis
1758:Mammals
1603:Bibcode
1522:1979296
1424:4854430
1389:6466405
1329:6670670
1233:2266605
1106:Bibcode
744:2825319
499:larvae.
430:-number
381:locusts
361:Odonata
357:Diptera
299:in the
237:mantles
171:insects
167:spiders
79:ocellus
67:circled
1763:horses
1725:Vision
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1009:572458
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396:cornea
383:) and
326:dorsal
319:oculus
297:f/0.58
280:dorsal
221:snails
206:cornea
194:, and
139:stemma
135:camera
131:retina
91:retina
1832:human
1753:Toads
1733:Birds
1467:S2CID
1385:S2CID
1283:S2CID
1187:S2CID
1005:S2CID
863:S2CID
832:(PDF)
821:(PDF)
698:4 May
569:pupal
527:fleas
349:wasps
313:Latin
262:This
225:slugs
219:Many
208:, or
182:Some
1794:Eyes
1773:cats
1768:dogs
1748:Fish
1647:ISBN
1619:PMID
1569:PMID
1536:link
1518:PMID
1459:ISSN
1420:PMID
1334:PMID
1275:PMID
1238:PMID
1179:PMID
1140:link
1122:PMID
941:ISBN
886:ISBN
784:PMID
749:PMID
700:2018
624:Egfr
413:The
345:ants
341:bees
276:wasp
223:and
210:lens
2074:Eye
1611:doi
1593:".
1561:doi
1508:doi
1451:doi
1412:doi
1377:doi
1373:122
1324:PMC
1316:doi
1267:doi
1263:193
1228:PMC
1220:doi
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1169:doi
1114:doi
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1036:doi
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739:PMC
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